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MYCOTAXON Volume LXXXIII, Pp

MYCOTAXON Volume LXXXIII, Pp

MYCOTAXON Volume LXXXIII, pp. 19-57 July-September 2002

PHYLOGENY OF : PARTIAL SYSTEMATICS SOLUTIONS FOR CORE OMPHALINOID GENERA IN THE (EUAGARlCS)

Scott A. Redhead Systematic and Eastern Cereal and Oilseed Research Centre Research Branch. Agriculture and Agri-food Ottawa. Ontario. Canada. KIA OC6

and

Fran90is Lutzoni, Jean-Marc Moncalvo, and Rytas Vilgalys Department ofBiology. Duke University Durham. NC 27708-0338. USA

Abstract: The of previously assigned to Ompha/ina sensu lato or is reevaluated in light ofrecent molecularly-based phylogenetic hypotheses. Nomenclatural complications involving generic and specific names, lectotypifications and changes to the Code are analysed. Lichenompha/ia gen. noy. (type hudsonianus, syn. Ompha/ina hudsoniana) is proposed for Jichenized fonner omphalinas. Ampulloc/itocybe gen. noy. (type Agaricus c1avipes, syn. ClilOcyhe c1avipes) is erected for its type species. is emended to include greyish species fonnerly included in , but excluding reddish brown species related to Ompha/ina pyxidata, the conserved lectotype for Omphalina. The genera , , Cerronema, Clabrocyphella, , , Hygrophorus, , Pseuduarmillariella, and , and the generic names Botrydina, Coriscium, Leptoglossum, Phaeotellus, Phytocunis, and Semiomphalina are discussed. 20 Key words: basidiolichen, Ampulloclitocybe,Arrhenia, , Omphalina, .

Introduction

As noted in the accompanying article on omphalinoid that may have evolved outside ofthe Agaricales (Redhead et ai, 2002), the omphalinoid and clitocyboid habits represent simple morphologies that presumably arose multiple times. This hypothesis is well-supported by analyses that distantly separate several omphalinoid clades, including a lineage encompassing Rickenella Raithelh., possibly outside ofthe Agaricales (Moncalvo et al. 2000 & 2002; Redhead et al. 2002). In the present paper, we discuss the systematics and biology of several genera with omphalinoid basidiomata, including clades containing the conserved lectotype for the name Omphalina Quel., viz. 0. pyxidata(Pers.: Fr.) Que!. (see Greuter et al. 2000, p. 192; see Kuyper 1995 for authorship), and type species for the names Arrhenia Fr., Leptoglossum P. Karst., and Phaeotellus Kuhner & Lamoure, as well as the lichenized omphalinoid taxa previously referred to a lichenized for which the rejected names Phytoconis Bory, Botrydina Breb., and Coriscium Vain. (see Greuter et al. 2000, pgs. 393-395), were applied. The discussion necessarily involves the generic name Gerronema Singer (1951 a), because Singer (1970, 1986), and those who followed his classification, included the lichenized omphalinoid taxa within the enlarged concept he ultimately adopted for that genus.

Stereotypical omphalinoid species fall into at least five inferred lineages within the greater Agaricales (= euagaric clade) using ITS and LSU data (Lutzoni 1997) and as supported by LSU data alone but using a much larger taxon sampling (Moncalvo et al. 2000 & 2002). These lineages are: (1) Gerronema sensu stricto (Singer 1951 a, emended by Redhead 1986) non Singer (1986), e.g., G. strombodes (Berk. & Mont.) Singer and G. subclavatum (Peck) Singer ex Redhead; (2) Chrysomphalina ClemenlYon sensu stricto (emended by Redhead 1986, Norvell et al. 1994) non Clemen~on (1982), e.g., C. chrysophylla (Fr.) ClemenlYon and C. grossula (Pers.) Norvell et al. (syn. Omphalina wynniae (Berk. & Br.) Ito); (3) the primarily bryophilous grey and brownish grey 2\ pigmented Arrhenia-Phaeo/ellus-Omphalina epichysium clade; (4) the lichenized omphalinas (Botrydina & Phytoconis of Redhead & Kuyper 1987, 1988, Norvell et al. 1994, Kuyper 1995); and (5) the group =Clitocybe sect. Pyxidatae sensu Bigelow (1974, 1985) and Lamoure (1974) pro par/e. A distinct omphalinoid lineage, (6) e.g., Rickenella and allies (Can/harellopsis Kuyper, Corylidia P. Karst., Redhead et aI., Redhead et aI., Sphagnomphalia Redhead et al.), which possibly evolved independently outside of the Agaricales (Moncalvo et a!. 2000 & 2002), is treated in a separate paper (Redhead et a!. 2002).

All 6 lineages were previously supported by the equally weighted parsimony and maximum likelihood analyses ofthe combined ITS and LSU data set (Lutzoni 1997, figs. 5a, 7). In larger taxon samplings (Moncalvo et al. 2000 & 2002), these clades of omphalinoid agarics often appear separated from each other and interspersed with clades of agarics and other fungi not previously analysed by Lutzoni (1997), some being additional omphalinoid agarics. We reproduce Lutzoni's (1997, fig. 7) as our Fig. 1, but with the various clades reclassified and renamed as discussed below.

Taxonomy and nomenclature

Gerronema was originally proposed for lignicolous tropical species (S inger 195 Ia) that were latter characterized, in part, by the presence of sarcodimitic tissues (Redhead 1986, Norvell et a!. 1994). Two such species, G. strombode.'l and G. .'lubclava/um, formed a monophyletic unit in the analyses by Lutzoni (1997). Subsequently they were shown to be distantly related to other omphalinas (Moncalvo et a!. 2000). In a much larger analysis (Moncalvo et al. 2002), these two species ofGerronema cluster with other taxa with sarcodimitic tissues (e.g., with species assigned to (Singer) Metrod, Hydropu.'l (KUhner) Singer, Megacollybia Kot!. & Pouz., cf. Redhead, 1987), which together form a well-supported lineage (to be treated in a separate paper). Hence, neither the lichenized omphalinas nor Rickenella and alljes are closely related to Gerronema sensu stricto. 22

1III••I1!1!1•••: Gertooema SlJtdava'u~' ] Gerronema 100 Gstrombcdes 'Hygxcybe' cilrinopallida ] "Hygrocybe' ~!I!!!!I.--~ g!"Jss~Ja • Chrysomp/lalina ] Chrysompha/ina cnryscpr.,1la Agaricaies ClJr;SWo:;i!iili11a A veiutlpeS A. epc.~ysium A sphagnicol. A. ~or.Olis _-- A. chlorocyanea Arrhenia Alobaia A awiscalpium 79 A. obs~urata A. griseopalljda 52 JIII.IIII!••• L. ve/urina 100 .c=7:L gnS€Ra L---""-----L.a1pma Uchenomphalia """'F,i:O---- L lJu·jscmana ----L. umbellifera op}'Xida!a ] o ,'i~ulicola Omphalina "CIilocyre'laterilia Spn,,;nompllaJia bre'libasidiata ....------LOIe;e.'c mc'C~;;n'iae RJckenella meilea hymenochaetoid clade 93 Rpse-JJOg'iseila ---- C:lI~umyces rosena MuftidiMJla coryJlC..idEs I(~M-:. .···········11!1100"v vema!is 1nucleotide substitution/100 s~es

Fig. 1. Most likely phylogenetic tree for Omphalinoid mushrooms based on a combined nuclear LSU and ITS rONA data set. Adapted from Lutzoni (1997, Fig. 7). Note the revised labelling ofthe clades proposed here and that "wynniae" and "grossula" are combined as C. grassula. 23 "Chrysomphalina" nom. invalid. (Haas,1962) was coined for two carotenoid forming taxa. The "type" C. chrysophylla, was retained in Chrysomphalina CI<~men90n ( 1982) when the name was validated, while the other, flaasiella venustissima (Fr.) Kot!. & Pouz.. was excluded and treated in the genus Haasiella Kotlaba & Pouzar (1966): see also Kost (1986) and Chiaffi & Surault (1996). Neither of the two presently recognized Haasiella species have been sequenced, but it seems possible that Chrysomphalina species might be their closest allies, and if nested among them. the name Haasiella would have priority. Following a detailed anatomical study, Kost (1986) however, concluded they were not closely allied.

When Chrysomphalina was val idated by Clemen~on (1982), he included two species. the type. C. chrysophylla, and Gerronema strombodes. while excluding ll. venustissima. As indicated above, G. stromhodes is unrelated to C. chrysophylla. Additionally, the European concept of"G. strombodes" represented a misapplication of that name. Gerronema strombodes is restricted to eastern North America. The European taxon known to Clemen

Chrysomphalina, as thus defined, appears to be a monophyletic taxon allied firstly to a sister genus Hygrophorus Fr. (Moncalvo et al. 2000 & 2002), and possibly allied to other hygrophoroid genera (Lutzoni 1997; Moncalvo et al. 2000 & 2002). The genus Hygrophorus sensu stricto (e.g., H. bakerensis A.H. Smith & Hesler, H. sordidus Peck) is clearly a distinct lineage and genus, distinguished by a presumed obligate ectomycorrhizal symbiosis, absence of carotenoids (pigmented by sundry other pigments, Gill & Steglich 1987). possession ofa different chemistry, and clamp connection formation. Chrysomphalina species have elongated basidia (Norvell et al. 1994) and because of the pachypodial subhymenium, a somewhat divergent lamellar trama, perhaps homologous to the divergent, slightly differently arranged lamellar tramas characterizing Hygrophorus sensu stricto. 24 The combined phylogenetic analyses by Lutzoni (1997) sometimes placed HJlgrocybe citrinopallida (A.H. Smith & Hesler) Kobayasi (another hygrophoroid taxon) in proximity to Chrysomphalina (with weak support). The analyses by Moncalvo et al. (2000, fig. 2, clade "K") sometimes shifted it closer (also with weak support) to the lichenized omphalinas ("phytoconis") and Arrhenia clade. With the addition of more genera, especially other hygrophoroid taxa, analysis of the most parsimonious trees found from heuristic searches (MP) and the bootstrap 50% majority rule consensus tree (B8) (Moncalvo et ai, 2002) continues to support the taxon pair Chrysomphalina-Hygrophorus. Hygrocyhe (Fr.) P. Kumm. (including its type H. conica (Schaeff.: fr.) P. Kumm.) appears to represent a monotypic lineage with two fundamental subgeneric divisions, e.g., H. (Schw.) Murr. & H. miniata (Fr.: Fr.) P. Kumm. [sect. Coccineae subsect. Squamulose], and H. !:>padicea (Fr.) P. Karst. & H. conica [sect. Hygrocybe subsect. Hygrocybe]. The surmised sister taxon to this monophyletic Hygrocybe clade is a well-supported clade (90% BS) with cyanophylla (Fr.) Redhead et al. and the wayward Hygrocybe citrinopallida, also known as Gliophorus citrinopallidus (A.H. Smith & Hesler) Kovalenko (1999). Chromosera Redhead, Ammirati & Norvell (1995) is a recently described monotypic genus established to account for a species alternately known as liIacifolia (Peck) A.H. Smith in North America and Omphalina cyanophylla (Fr.) QueJ. in Europe. This lignicolous species was clearly neither an Omphalina nor a Mycena (Pers.) Roussel, but interestingly enough, KUhner (1980) suggested that it belonged in Hygrocybe subg. Gliophorus, and called it "Hygrocybe" cyanophylla nom. inval. (see Redhead et al. 1995).

Both Gliophorus Herink and Chromosera are characterized by viscid stipes and pilei, and by bright pigments. Gliophorus species are terrestrial and their biological status remains uncertain. Another species, G. laetus (Pers.) Herink, appears on a largely omphalinoid clade (the Phytoconis, Arrhenia, and Cantharellula- lineage, discussed below), i.e., separate from G. citrinopallidus. Unfortunately, as the type for Gliophorus, G. psittacinus (Schaeff. Fr.) Herink, has not been included in any ofthe analyses, application ofthe generic name is unresolved. The name Gliophorus potentially could compete with Chromosera, or be applicable to a sister taxon. 25 is a terrestrial species with an unestablished biological role (?ectomycorrhizal). In the analyses by Moncalvo et al. (2000, 2002), there is weak bootstrap support linking the Chromosera-Hygrocybe clade to the Chrysomphalina-Hygrophorus clade together with Humidicutis (Singer) Singer, a third hygrophoroid genus. Bootstrap analysis (but not MP analyses) support the phylogenetic placement of Humidicutis marginata (Peck) Singer (type for that generic name) next to the Chrysomphalina-Hygrophorus clade. These results potentially bring into inferred evolutionary proximity, a group of taxa that form muscaflavin based pigments (as surmised from other chemically analyzed Hygrocyhe) and other Hygrophorus, known to be able "to induce extradiol cleavage of L-DOPA leading to the formation of muscaflavin" (Gill & Steglich 1987, p. 86). Humidicutis is in part defined by the absence ofclamp connections and is otherwise different because of pigment characters (see Singer 1986). There are some indications that the core of the hygrophoroid taxa (the classical ""), are basal to the Agaricales (euagaric clade). In the analyses ofa portion ofthe mt LSU rRNA gene by Bruns et at. (1998), representatives of Hygrophorus and Hygrocyhe actually appeared outside of the depicted main agaric clade, whereas in the analyses by Hibbett et at. (2000), Hygrocyhe conica, Humidicutis marginata (as Hygrocyhe marginata (Peck) Murr.) and the two Hygrophorus species analysed by Moncalvo et al. (2000) were shown to be basal to the rest of the euagaric clade. In all analyses, the backbones of all resulting phylogenetic trees are weakly supported, except for those using fewer taxa combined with several genomic regions evolving at different rates (ITS region combined with LSU nuc rONA in Lutzoni 1997). We regard the Hygrophoraceae s.1. and s.s. (whether or not recognized as a distinct taxon) to be among the Agaricales (euagarics). Future investigation of that traditional family should include Chrysomphalina, Chromosera, possibly Haasiella, and the two omphalinoid lineages "Phytoconis" (= Lichenomphalia, see below) and Arrhenia as emended below.

The primary goal ofthe study by Lutzoni (1997) was to investigate the phylogeny of lichenized omphalinas and their relationships to non­ -forming omphalinoid agarics. Fourofthe five sampled lichenized species (Omphalina grisella (P. Karst.) Moser, 0. hudsoniana (Jenn.) Bigelow,o. luteovitellina (Pilat & Nannf.) M. Lange, and 0. velutina 26 (Quel.) Quel.) formed a clearly defined, strongly supported (BS = 92­ 98%) monophyletic lineage. The fifth, 0. ericetorum (Pers.: Fr.) Bigelow (see discussion on authorship below and synonymy with "umbellifera"), was inferred to be sister to this lineage in all three combined analyses of the ITS region with the LSU nuc rONA, but the placement received weak support (BS < 50-70%; Lutzoni 1997: figs. 5 & 7). The sister group to the lichenized clade in the 1997 analyses was a nonlichenized, largely bryophilous group of omphalinoid-arrhenoid species, all with dark pigments. This sister relationship was supported by bootstrap support ranging from 70-83% when the ITS region was combined with LSU data (Lutzoni 1997). Deeply nested among the lamellate omphalinoid taxa were the reduced agarics, Arrhenia auriscalpium Fr. (type for Arrhenia), (Pers.: Fr.) Redhead (type for Dictyolus Quel.), and Phaeotellus griseopallidus (Desm.: Fr.) Kuhner & Lamoure. In a larger taxon sampling restricted to the LSU data (Moncalvo et al. 2000), this close relationship between lichen-forming omphalinas and the grey and grey-brown omphalinoid­ arrhenoid clade continues to be detectable with even larger datasets (Moncalvo et al. 2002), but the exact relationship ofthe taxon labelled 0. ericetorum (= umbellifera) became uncertain with regard to these two lineages because of the paucity of characters inherent to a single gene phylogenetic analysis for such a large taxon sampling. In the smaller taxon matrix, a third bryophilous omphalinoid clade identified by Lutzoni (1997; BS = 95-100%, Figs. 5 and 7) and labelled "Clitocybe" (i.e., Omphalina pyxidata, 0. rivulico/a (1. Favre) Lamoure, Clitocybe /ateritia J. Favre) appeared to be a sister taxon to the "Omphalina"­ Chrysomphalina clade, with bootstrap support in all combined analyses ranging from 52-83% (Lutzoni 1997, Figs. 5 and 7). However, support weakened when diluted by the addition of far more diverse taxa and when phylogenetic analyses were restricted to LSU (Moncalvo ct al. 2000). The 0. pyxidata-C. /ateritia clade (in clade "J") no longer groups with any other identifiable former Omphalina. MP analysis using a far larger taxon sampling suggests a relationship both to the bryophilous genus Rimbachia Pat. (R. bryophi/a (Pers.: Fr.) Redhead) and (less closely) to "Clitocybe" clavipes (Pers.: Fr. ) P. Kumm. (a taxon which remained distantly located in Moncalvo et al. 2000, fig. 2). None of these were close to the core Clitocybe (Fr.) Staude (eg. C. nebularis (Batsch: Fr.) P. Kumm., C. dealbata (Fr.: Fr.) P. Kumm., Lepista nuda 27 (Fr.: Fr.) Cooke), which appear to be far more closely related to the former Lyophylleae (Hofstetter et ai, in prep.).

The conserved lectotype for the generic name Omphalina Que!. is 0. pyxidata (Greuter et al. 2000, p. 192), hence we will hereafter apply the generic name Omphalina to the presumably bryophilous clade centered around the type, 0. pyxidata, and not refer to it as "Clitocyhe" as it was tentatively labelled with a question mark by Lutzoni in 1997. Conservation of the lectotype for Omphalina ends decades of debate (Bigelow 1970, Donk 1962, Jorgensen & Ryman 1989, 1994, Lange 1981, Redhead & Weresub 1978, Redhead 1993, Redhead & Kuyper 1987, 1988, 1993, Singer 1962, 1975, 1986, Singer & Smith 1946), but with unanticipated repercussions. As thus defined and restricted, Omphalina is not congeneric with either the lichenized species or the grey-brown nonlichenized "omphalinas". Biologically, Omphalina (sensu conserved lectotype) occupies similar habitats and substrates as species in the Arrhenia lineage, but they differ in subtle pigmentation chemistry and topology (the presumed melanins are more reddish brown than grey brown, and the deeper tissues are pigmentless, i.e. whitish). Without sequence data as a guide, these differences would not normally be considered to be generically significant, especially when compared to far more conspicuous morphological differences. Nonetheless, they do appear to correlate with base-pair sequence data generated phylogenies.

A identified as Omphalina pyxidata has been linked by Hornby & Ward (1995) to a sclerotium-forming, weak pathogen of turf grass roots. However, they reported unspecified genetic differences between herbarium specimens of 0. pyxidata and the suspected causal agent; specimens were not retained (Hornby 1995, pers.com.). Additionally the pinkish-purple coloration of the basidiomata (illustrated in color) suggests that another fungus might be involved. Therefore, its generic identification remains ambiguous.

Notwithstanding the above report, Omphalina pyxidata has been regarded as a bryophilous fungus (Bigelow 1974, Lamoure 1974). Similarly 0. rivulicola is bryophilous (Lamoure 1974). Together they form a bryophilous clade (Moncalvo et aI., 2002). Sister to this 28 bryophilous clade was one containing an unidentified Clitocyhe sp. and Clitocybe lateritia. The latter grows with Dryas litter in alpine meadows (Lamoure 1972) and presumably is not bryophilous. In common with 0. pyxidata and 0. rivulicola, C. lalerilia produces reddish brown melanin­ like pigments, rather than the greyish pigments of the Arrhenia clade. We will return to the discussion ofthe Omphalina (0. pyxidata clade) further below.

The oldest available generic name for any of the other omphalinoid species that appear on the combined Arrhenia-"Phytoconis" clades is Arrhenia Fries (1849). Arrhenia was first erected for basidiomycetes characterized by nutant basidiomata with cantharelloid hymenia (Comer 1966, Heiland 1976, Pilat & Nannfeldt 1954). Later the genus was expanded to include species with pleurotoid basidiomata, incl. Leploglossum and Phaeotellus (Aronsen 1992, Breitenbach & Kranzlin 1991, Courtecuisse 1986, GuIden 1987, Gulden & Torkelsen 1996, Kreisel 1987, Kuhner & Lamoure 1972, Moser & Julich 1997, Redhead 1984, Senn-Irlet 1987, Watling 1988, Watling & Gregory 1989). However, the delimitation from omphalinoid taxa has always been controversial (Gulden 1987, Heiland 1982, Lange 1981, Watling 1988), as has been the delimitation of Arrhenia from Leptoglossum (Corner 1966, Redhead 1984), including debatably classified species with centrally stipitate basidiomata, and cantharelloid hymenia. There is a complete transitional series of morphological features from gross omphalinoid to cyphelloid habit, and lamellate to merulioid to smooth hymenia. Based upon the deeply nested position ofArrhenia within the clade ofmore typically omphalinoid taxa (Lutzoni 1997, Moncalvo et al. 2000 & 2002) and the gradation of morphological features, we are now prepared to expand the coverage of the name Arrhenia to include omphalinoid, pleurotoid, nutant, and cyphelloid forms to this primarily bryophilous/phycophilous lineage with greyish melanized basidiomata. Invasion ofdead cells has been documented (Hassel & Kost 1998), at least in the instance ofthe species with the most reduced basidiomata, A. reliruga (Bull.: Fr.) Redhead, a cyphelloid taxon and apparent necrotrophic parasite. Although described on old bovine dung, the grey­ brown, c1ampless, cyphelloid fungus, Glabrocyphella stercoraria Barrasa, Esteve-Rav. & Sanchez (Barrasa et al. 1998), is best considered to be a species of Arrhenia perhaps associated with the algae also 29 reportedly growing on the dung substrate. Notably, the possibility of microscopic moss protonema was not excluded. For now we are prepared to include the similar species, Leptoglossum omnivorum Agerer (1984), in Arrhenia, based in part on exam ination of the type. These cyphelloid and nutant taxa are morphologically term inal groups of the primarily dark grey "omphalinoid" group ofspecies (cf. Lamoure 1975, Singer 1956).

The taxonomic question ofwhether or not to distinguish the lichenized "Phytoconis" generically from nonlichenized Arrhenia hinges in part on conveyance of information regarding phylogeny, biological status, and anatomy, and in part on nomenclature. It has previously been argued based upon anatomy and biology that the lichenized omphalinas should be recognized in a distinct genus (Kuyper 1986, 1994, 1995, Norvell et al. 1994, Redhead & Kuyper 1987, 1988). Lutzoni & Vilgalys (1995a) confirmed these results (BS = 75%, Figs. 2A) with a phylogenetic analysis restricted to morphological and anatomical characters. When combined with LSU data, the bootstrap value dropped slightly to 67% (Lutzoni & Vilgalys 1995a, Fig. 7). In the phylogenetic study with the most molecular characters (i.e., ITS region combined with LSU nuc rONA, Figs. 5 and 7, Lutzoni 1997) the three combined analyses consistently revealed the lichenized omphalinas as a monophyletic entity with bootstrap support as high as 70%. Other studies with broader taxon sampling but restricted to the LSU (Moncalvo et al. 2000, & 2002) did not have sufficient data to provide bootstrap values « 50%) for the phylogenetic placement of "0mphalina" ericetorumlumbellifera specifically on this clade; rather it is only placed in proximity. The latter studies do not confirm nor contradict the monophyly of the lichenized omphalinas. Therefore, our best phylogenetic estimates (i.e., those with the highest level ofconfidence) come from the phylogenetic analyses of morphological data alone (BS = 75%; Lutzoni & Vilgalys, 1995a) and from the phylogenetic analyses with the most molecular data, i.e., combined ITS with LSU data set (BS up to 70%; Lutzoni 1997). Both analyses independently supported the monophyly ofthe lichen-forming omphalinas. None ofthe competing phylogenetic relationships for the paraphyly or polyphyly ofthe lichenized omphalinas received bootstrap support > 50%. We therefore opt to recognize the lichenized omphalinoid agarics as a distinct genus. 30 Unfortunately, all three applicable validly published generic names ­ Phytoconis, Botrydina. and Coriscium - have been placed on a list of rejected names (see Gams 1995; Greuter et al. 2000) and hence are unavailable. This regrettable nomenclatural consequence results from not applying the same standards to basidiolichen generic names as is done for ascolichen generic names. To quote from the report of the Committee for Fungi (Gams 1995, p. 413), "By consequence those accepting the lichenized species as a separate genus are instructed to describe and name it as new, based upon holomorphic material, rather than using the anamorph name Phytoconis which was taken up by Redhead & Kuyper (in Mycotaxon 31: 321-323.1988), following the letter (but not the intention) of Art. 59." Therefore, despite the recognition ofa distinct wholly lichenized genus and acceptance ofthe name "Phyloconis" by agaricologists (ClemenlYon 1997, Collin & Lauron 1994, Grgurinovic 1997, Kuyper 1995, Nezdoiminogo 1997, Norvell et al. 1994, Redhead & Kuyper 1987, 1988, Romagnesi 1992, Watling 1988), the name Phytoconis is no longer available. We therefore name a "new" genus, Lichenomphalia gen. nov. (see below).

Lichenomphalia appears to have evolved from a bryophilous or phycophilous ancestor as suggested by Redhead & Kuyper (1987) and confirmed by Kranner & Lutzoni (1999). Using Omphalina as a model system (cf. Lutzoni & Vilgalys 1995a & b, Lutzoni 1997, Lutzoni & Pagel 1997), Kranner & Lutzoni (1999) noted that among analyzed Lichenomphalia (as lichenized Omphalina) and the non-I ichenized sister genus Arrhenia (as Arrhenia, Omphalina, and Phaeotellus), L. umhellifera comb. nov., Arrhenia velutipes comb. nov., and A. epichysium comb. nov. (see below), are the most basal and exhibit the slowest evolutionary rate changes in their respective clade. Therefore, they are the closest living phenotypic representations of the common ancestor that the non-lichenized Arrhenia and lichenized Lichenomphalia share. All three species are bryophilous and/or phycophilous. Kranner & Lutzoni (1999, Fig. I) identified three major predispositions for a successful transition to a lichenized state: I) high phenotypic plasticity/broad ecological amplitude, 2) low fungal virulence/high photobiont infection resistance, 3) desiccation and sun irradiation tolerant/efficient DNA repair mechanisms. They further noted that L. umbellifera (as 0. ericetorum) is the most plastic species ofthe 31 lichenized Omphalina group, as well as the most common, the most broadly distributed, and with the broadest range of substrates. This fulfils the first requirement. The study of predacious fungi lysing colonies of algae, bacteria, pollen, and/or yeasts (Barron 1988, Hutchinson & Barron 1996, 1997a, b) using modified hyphae remarkably similar to those forming the thalli of these and other basidiolichens (Oberwinkler 1970, Redhead & Kuyper 1987), suggests a physical mechanism and chemo-attractant response to algal colonies that provided an opportunity for the lichenization process to begin. It has also been shown that icmadophilae Jaag, the alga entrapped by Lichenomphalia, forms cell walls resistant to Iysolytic enzymes (Honegger & Brunner 1981). Thus, the combined low-level parasitism and resistance to enzymatic lysing fulfils the second requirement.

Lutzoni & Pagel (1997) outlined phenomena supporting fulfilment ofthe third requirement, i.e., strong positive selection in the lichenized omphalinas toward replacing one of two adjacent thymines by another nucleotide to prevent thymine dimerization. Given the poikilohydric and heliophylic nature of most , lineages tolerant to oxidative stress caused by desiccation and sun irradiation, and/or with efficient DNA repair mechanisms would be more suited to forming a successful mutualistic association with terrestrial cyanobacteria or green algae. Again, L. umbell(fera appears to exhibit more ancestral features because its thallic mycelium is broader and not as thick-walled (Redhead & Kuyper 1987, as Botrydina botryoides (L.) Redhead & Kuyper) and therefore, presumably less tolerant ofradiation and dehydration damage, it occupies wetter habitats on average, and its basidiomata are less protected by strong pigments (either dark melanins or the bright yellow­ orange uncharacterized pigments).

We anticipate that the as-yet unsequenced and rare genus, Semiomphalina Redhead (1984), a nutant, pale arrhenioid but lichenized basidiomycete (known only from the type collection of S. !eptog!ossoides (Corner) Redhead) will prove to be a sister taxon to Lichenomphalia. Its and thallus resemble those ofL. umbellifera. 32 Other omphalinoid/clitocyboid genera

In both the final BS and MP trees generated for the Agaricales analyses (Moncalvo et al. 2002), a moderately supported clade (49% BS) bearing (J.F. Gmelin: Fr.) Singer and Pseudoarmillariella ectypoides (Peck) Singer (types oftheir respective generic names) appears together on a weakly supported clade bear.ing the Arrhenia clade (77% BS) and all sampled Lichenomphalia as defined above (L. velutina, L. alpina, L. hudwniana - on one clade with 77% BS; and L. umbellifera separately), and also . Anatomically, the Cantharellula-Pseudoarmillariella clade is generically easily distinguished from nonamyloid-spored Arrhenia, Lichenomphalia, and Gliophorus laetus by the formation of amyloid spores.

Pseudoarmillariella ectypoides is lignicolous (a white rot agent?) and has a scaly, orangish brown pileus. Cantharellula umbonata is primarily bryophilous, but occasionally remains in residual populations in young conifer forests with recently, closed canopies where populations ofhost have died (Redhead & Malloch 1986). Species in both Pseudoarmillariella and Cantharellula share characters such as forked lamellae, reddish brown bruising reactions of the hymenium, and similarly arranged subhymenial elements, i.e., "subirregularly intermixed-subramose" elements "strongly interlaced-curved in all directions and therefore at times appearing cellular" (Singer 1986 [when comparing Pseudoarmillariella to Cantharellula]). Previously Singer (1951 b) had treated both type species in one genus, Cantharellula. The generic name Pseudoarmillariella (Singer) Singer was in fact an elevation in status of Cantharellula subg. Pseudoarmillariella Singer. Interestingly, Bigelow (1982) had transferred P. ectypoides to Omphalina sensu Bigelow (ca. = Gerronema sensu Singer), but he maintained the distinction between it and Cantharellula (Bigelow 1975). We maintain the separate generic identities for Pseudoarmillariella and Cantharellula as proposed by Singer (1956) and outlined by Singer (1986), while still acknowledging the close relationship between them. As noted by Norvell et al. (1994), Pseudoarmillariella possesses two pigment classes and forms a thickening hymenium, in addition to being a wood decay fungus. 33 Clilocybe c1avipes, which occurs on a weakly supported clade with Rimbachia and Omphalina sensu lectotype (Moncalvo et ai, 2002, fig. 2), presents a dilemma because it seems to represent a distinct genus, phylogenetically more closely related to the type of Omphalina than the type ofClilocybe (see discussion below). The sequence data arc unlike other traditional clitocyboid taxa, most of which prove to be more closely related to the Lyophylleae (Moncalvo et al. 2002; Hofstetter et al. in prep.). Clilocybe cfavipes is uniquely characterized among c1itocyboid taxa by the formation ofcoprine-like compounds (Cochran & Cochran 1978, Yamaura et a1.1986) and by ultrastructurally minutely roughened basidiospores (Pegler & Young 1971, Bigelow 1981, 1982) compared to smooth-spored Clilocybe or echinate-spored Lepisla (Fr.) W.G. Smith. Clilocybe c1avipes also produces a novel class oftyrosine kinase inhibitors, named the c1avilactones (Cassinelli et al. 2000). The combination of significantly different sequence data from other "Clilocybe", and ultrastructurally distinctive spores suggest that "e. cfavipes" has been incorrectly classified as a Clilocybe. We therefore propose to recognize a distinct genus, Ampuflocfitocybe (type = A. cfavipes, see below) for the species. Strict consensus ofthe 5,000 most parsimonious trees found (MPF) by Moncalvo et al. (2002) place Ampuffoclitocybe c1avipes next to Rimbachia bryophila, which in turn is sister to the Omphalina (0. pyxidata) and Clitocybe laleritia clades.

The phylogenetic proximity ofthe fungus, Ampuffoclitocybe cfavipes to a fungus named Clitocybe lateritia (Moncalvo et al. 2002), and the phylogenetically distanttaxa named Clilocybe dealbata, C. odora (Bull.: Fr.) P. Kumm., e. ramigena Bigelow, e. connata (Schum.:Fr.) Gillet (=Lyophyffum connatum (Schum.: Fr.) Singer), e. glacialis Redhead ct al. (=Lyophyffum montanum A.H. Smith), C. nebularis (=Lepista nebularis (Batsch: Fr.) Harmaja), and e. nuda (Fr.: Fr.) Bigelow & A.H. Smith (=Lepista nuda), begs a question regarding application of the name C/ilocybe (Fr.) Staude and its lectotypification. The lectotypification of Cliloc-ybe has been as heatedly debated as has lectotypification of Ompha/ina, and the topic remains unsettled (Bigelow 1965, Clements & Shear 1931, Donk 1949, 1962, Harmaja 1969, 1974, Horak 1968, Imai 1938, Korf 1988, KUhner 1980, Kuyp~r 1982, 1995, Rauschert 1986, Singer 1936, 1951 b, 1962, 1975, 1986, Singer & Smith 1946). The choice of C. c1avipes (type of our 34 Ampulloclitocyhe) as lectotype of Clitocybe by Bigelow (1965) is noteworthy, but must be dismissed because of the earlier lectotypifications (Greuter et al. 2000, Art. 9.17). The primary division of thought is split between support for Agaricus nebularis Batsch: Fr. versus Agaricus infundihuliformis Schaeff. (=Agaricus gibbus Pers.: Fr. according to Singer 1986; = Agaricus flaccidus Sow.: Fr. according to Fries 1821 and Kuyper 1995). Donk (1962) reviewed the various typifications proposed up to 1962. The earliest lectotypification of the generic level name "ClitoGybe" [with variously attributed authorship all involving Fries and his "tribus" as basionym] choosingA. nebularis, was by Earle (1909) followed by Murrill (1916), both following the American Code of Nomenclature. Such choices made using the American Code may be superseded (Greuter et al. 2000, Art. 10.5b). Using a misspelled epithet with incorrect, non-existent authority, Clements & Shear (1931) selected "e. infundibulis (Schaeff.) Fr." Despite the errors, their intent was clear, but because there is, in fact, no such species, their lectotypification [for priority purposes] is rejected. Singer (1936) and Singer & Smith (1946) were the next to name a [lecto­ ]type, namely ClitoGybe infundibuliformis, but for either Clitocyhe "Fr." or Clitocybe "(Fr.) Quel." However, the names Agaricus in.fundihuliformis Schaeff. and A. infundibuliformis Bull., were both treated as synonyms ofAgaricusflaccidus Sow.:Fr. (an accepted, named species) by Fries (1821). Technically the choice ofA. in.fundibuliformis is rejected simply because in the protologue of the basionym for the generic name, i.e. Agaricus "tribus" Clitocybe Fr. (1821: 78), that particular species binomial was not used for a named, recognized species. The first choice ofa recognised, named species - both in Fries' "tribus" and when elevated by generic status (by Staude) - was Donk's (1949) selection of A. nebularis. Nonetheless, unless conserved, lectotypification of Clitocybe (Fr.) Staude will remain debatable. A proposal to conserve ClitoGyhe (Fr.) Staude with a conserved lectotype, Agaricus gibbus Pers.: Fr., by Rauschert (1986) was rejected (Korf 1988) simply because the proposal lacked detail, and unfortunately the proposer had subsequently died. For our purposes, we accept e. nehularis (cf. also Kuyper, 1992). The generic taxonomic delimitations (Clitocybe versus Lepista and various Lyophylleae) and precise phylogenetic placement ofClitocyhe (typified bye. nebularis), which is phylogenetically distant from the clade with Omphalina (0. pyxidata), 35 "Clitocybe" lateritia, and Ampulloclitocybe c1a\'ipes, as well as the clade with Arrheniaand Lichenomphalia, are the subjects ofother manuscripts in this series. At this time we lack sufficient confidence and information to assign a different generic name to Clitocybe lateritia. Possibly it will be assigned to the reconstructed genus Omphalina.

For an interim key to the omphalinoid genera discussed above we refer to the key in Norvell et al. (1994), replacing the name "Phytoconis" with Lichenomphalia, and replacing couplet #8 (leading to Omphalina, Clitocybe) with the following couplets.

8. Bryophilous or phycophilous (sometimes appearing terrestrial or lignicolous due to sparse growth of associated algae or bryophytes), never on sound clean wood or conspicuously causing white or brown wood rots [with incrusting pigments, odor never farinaceous]; reduced forms exhibit hymenial folds and wrinkles, may lack stipes. may be nutant or pendant [see also key to Arrhenia by Redhead, 1984] .. .. 9 8. Terrestrial or lignicolous, not dependent on living algae or bryophytes; ifon wood, mycelium clearly penetrating and decaying wood [easily cultured on standard general media from spores, odor sometimes farinaceous or strongly aromatic, many species large and not omphalinoid] 10 9. Coloration of pilei and stipes grey, blackish, or bluish, and brown­ grey, with slightly paler but similarly colored hymenia (lamellae or folds), or for some sphagnicolous species pilei scaly centrally with darkly incrusted pigments but pileus tending to be beige or tan and then lamellae are pallid; clamps present or absent, stipes and/or regular lamellae present or absent Arrhenia 9. Coloration of pilei and stipes reddish brown (rusty, vinaceous brown, orangish brown) and lamellae contrastingly much paler (i.e. pallid, whitish, pastel), always smooth and not scaly; clamp connections present; lamellae always well-formed Omphalina 10. Basidiomata nonhygrophanous, with markedly darkened fleshy pileus and fleshy bulbose-based stipe contrasting with the whitish cream lamellae; clamps present; pigments intracellular; spores smooth (light microscopy) but ultrastructurally rugose (SEM), not cyanophilous or congophilous, white in prints; producing coprine-like pOisonmgs 36

...... Ampu/loclitocybe 10. Not with above combination other unresolved Clitocyboid genera.

Note: It should be recognized that the reclassification based upon phylogenetic studies is a work in progress.

New genera and new combinations

Ampu/loclitocybe gen .noY. Basidiomata clitocybijormis. Non lichenisatus. Trama lamellarum bidirectionalis, hyphis fihulatis. cystidia nulla. Sporis alhis, microrugosis, acyanophilis, inamyloideis. Type: Ampu/loclitocybe clavipes (Pers.: Fr.) comb. noy. Basionym: Agaricus clavipes Pers., Syn. meth. Fung. p. 353. 180 I. (A. clavipes Pers.: Fr., Syst. mycol. I: 86. 1821)

Lichenomphalitl gen noy. Basidiomata omphalijormis. Lichenisatus. Trama lamellarum bidirectionalis, hyphis defibulatis, cystidia nulla. Sporis alMs, inamyloideis. Type: L. hudsoniana (see below). = Phytoconis Bory 1797 nom. rej. (Greuter et aJ. 2000) =Botrydina Breb. 1839 nom. rej. (Greuter et al. 2000) = Coriscium Vain. 1890 nom. rej. (Greuter et al. 2000) = Omphalina sect. Defibulatae Singer 1975 = Gerronema sect. Phycophila ClemenIYon 1982 =Clitocybe subsect. Defibulatae (Singer) Bigelow 1985

Lichenompha/ia species

L. alpina (Britzelmayr) comb. noy. Basionym: Agaricus alpinus Britz., Ber. Naturw. Ver. Augsburg 30: 13. 1890.

Comments: Synonymous names are: Cantharellus dovrejjeldiensis P. Henn. & Kirschst., Agaricus umbellijerus var. flavus Sommerf., Omphalia flava (Sommerf.) Moller, Omphalia luteovitellina Pilat & Nannfeldt (see Jergensen & Ryman 1989b). 37 Application and synonym of the name Agaricus a/pinus has been disputed (Redhead & Kuyper 1987, Jorgensen & Ryman 1989b). The designation of Britzelmayr's plate by J0rgensen & Ryman (1989b) as lectotype did little to resolve the controversy. Therefore, we hereby designate an epitype for the name Agaricus a/pinus Britz., i.e. the type for Ompllalia luteovitellina. If we do not do so, thereby fixing application of the name to this taxon, then we are forced to adopt Cantharellus dovrefJe/diensis as the next available name (J0rgensen & Ryman 1989b). We cannot fully explain the subtle discrepancies, such as paler stipe, and perhaps slightly hollowed stipes recorded by Britzelmayr (1890), neither of which is normal for the taxon accepted here. Neither discrepancy appeared to bother J0rgensen & Ryman ( 1989b), nor the fact the species as accepted here has not been independently documented in the region where Britzelmayr collected in the past century. However, Favre (1955) found this species, reported as Omphaliajlava, in alpine regions of the Swiss Alps, and therefore it is possible that Britzelmayr had found it in the somewhat nearby Bavarian alpine regions. The synonym proposed by Redhead & Kuyper (1987), i.e. placement ofAgaricus a/pinus Britz. in synonymy with Endocarpon viride Ach. (= Botrydina viridis (Ach.) Redhead & Kuyper = Lichenomphalia hudsoniana), is rescinded simply for convenience and to reach consensus with J0rgensen & Ryman (1989b).

The report ofOmphalina a/pina (Britzelmayr) Bresinsky & Stangl (with Omphalia /uteovitellina Pilat & Nannfeldt (1954) in synonymy) from Yunnan was based upon a fungus reportedly with clamp connections (Horak 1987: 179). If the presence of clamp connections is true, that fungus cannot be conspecific with any Lichenomphalia.

L. aurantiaca (Redhead & Kuyper) comb. nov. Basionym: Botrydina aurantiaca Redhead & Kuyper, Arctic & Alpine Mycology II, p. 334. 1987. Plenum Press, NY.

L. cllromacea (Cleland) comb. nov. Basionym: Omphalia chromacea, Trans. & Proc. Roy. Soc. South Australia 48: 239. 1924.

Comments: An incorrect basionym citation was given by Redhead & Kuyper (1987, 1988) and corrected by May & Wood (1995). Isotype 3X material at BPI was recorded by Redhead & Kuyper (1988).

L. grisella (P. Karst.) comb. nov. Basionym: Omphalia grisella P. Karst., Medd. Soc. Fauna Flora Fennica 16: 92. 1890 [ut "(Weinm.?) n. Sp."].

L. hudsoniana (Jennings) comb. nov. Basionym: Hygrophorus hudsonianus Jennings, Mem. Carnegie Mus. III, 12: 2. 1936.

Comments: Synonymous names are: Endocarpon viride Acharius [thallus only, rejection ofthis name has been recommended - cf. Gams (1995), and is somewhat listed as such in Appendix IV of the Code (Greuter et aI., 2000: 394) as type for the rejected generic name Coriscium Vain.], Verrucaria laetevirens Borrer [thallus only], Omphalia luteolilacina Favre (fide Redhead & Kuyper 1987), but not Agaricus alpinus Britz.(see discussion above).

L. lobata (Redhead & Kuyper) comb. nov. Basionym: Botrydina toba/a Redhead & Kuyper, Arctic & Alpine Mycology II, p. 334. 1987. Plenum Press, NY.

L. umbellijertl (L.: Fr.) comb. nov. Basionym: Agaricus umbelliferus L., Species plantarum p. 1175. 1753; Agaricus umbelliferus L.: Fr., Elenchus fungorum, p. 22. 1828.

Comments: Synonymous names are: Byssus botryoides L. [thallus only­ see Redhead & Kuyper 1987, Jorgensen et al. 1994], Botrydina vulgaris Breb. [thallus], Agaricus pseudoandrosaceus Bull., Agaricus nothus Gmelin, Agaricus ericetorum Pers.: Fr., Agaricus valgus Holmskjold, Merulius turfosus Pers., Omphalia luteola Peck, Omphalia sphagnophila Peck, Omphalinajulvopallens P.O. Orton (see Redhead & Kuyper 1987, 1988, Kuyper 1995, except for their discussion on A. umbelliferus as is detailed below).

Following the extensive review of names and types discussed by Redhead & Kuyper (1987), Jorgensen & Ryman (1989a & b, 1994) published nomenclatural reviews of their own and designated several neo- or lectotypes, some placing names in conflict with those adopted or 39 rejected by Redhead & Kuyper (1987), and others supporting the interpretations by Redhead & Kuyper (1987, 1988). The most contentious and destabilizing conflict revolves around the choice of Agaricus ericetorum Pers.: Fr. versus Agaricus umbelliferus L.: Fr. Jorgensen & Ryman (1989b) incorrectly claimed (i) that Redhead & Kuyper (1987) had overlooked the fact that Fries (1821) "only cites his own Obs. mycol. I p. 87" and (il) that Fries did only that. In fact Fries (1821: 509) specifically gives the authority as "ericetorum P. A. S." not "Fr." as he did for his own named species ( "P." = Persoon and "A.S." = Albertini & Schweinitz). These facts were explicitly noted by Redhead & Kuyper (1987: 326-327). Notwithstanding all previous arguments published, we note that in the latest Code (Greuter et al. 2000: 30, Art. 15.1 Ex. I) the two names are cited as "Agaricus ericetorum Fr." and "A. umbelliferus L." which suggests that "Agaricus ericetorum Fr.: Fr." is a quasi-conserved (i.e. unlisted in any appendix ofthe Code) name with that authority, rather than "Agaricus ericetorum Pers.: Fr." This makes the citation of"Agaricus ericetorum Fr." in the Code an ambiguous and poor example. We further note that changes in articles and wording of the Code (Greuter et al. 2000) again alter nomenclatural conclusions reached using earlier Codes by various authors. Specifically, Art. 7.5 now states, "Automatic typification does not apply to names sanctioned under Art. 15." Hence, Agaricus ericetorum of Fries (1821) would not be automatically typified by the type ofA. pseudoandrosaceus Bull., as it would have been under the Code in effect (Voss et al. 1983) when Redhead & Kuyper (1987) interpreted the names.

What, then, are the lectotypes for the names Agaricus umbelliferus and Agaricus ericetorum, how many names with different authorities are there, and - as ultimately typified - are they taxonomic synonyms? Or in other words, how many homonymic names called Agaricus ericetorum are there, and which is sanctioned by Fries' (1821) usage of that binomial?

Agaricus ericetorum Pers. (1796) was first introduced as a binomial citing the earlier named Agaricus pseudo-androsaceus Bull. pI. 276 (Bulliard 1786) in synonymy. Agaricus ericetorum Pers. is therefore illegitimate (Art. 52.1), unless it is considered sanctioned (Art. 15) or conserved (Art. 14). The concepts ofboth Bulliard (1786) and Persoon 40 (1796) were broad and as currently recognized included more than one species. Redhead & Kuyper (1987: 329) lectotypified Agaricus ericetorum Pers. and Agaricuspseudo-androsaceus Bull. by designating the same illustration on Bulliard plate no. 276, specifically choosing the central figure of pale, rather than greyish agarics depicted on a moss cushion, and thus exclude the greyish forms. This typification of Agaricus pseudo-androsaceus Bull. was uncontested and accepted by Jorgensen & Ryman (1989a: 140). All authors agreed it was applicable to the species represented by Lundell & Nannfeldt, Fungi exsicc. Suec. No. 1753.

Singer (1961: 23) had previously selected a specimen ofA. ericetorum from Persoon's herbarium in Leiden (L91 0.256-1888) as "lectotype" for Agaricus ericetorum "Pers. ex Fr.", but it can at best be considered to be an epitype (or neotype) because there is no date on the specimen and it was not seen by Bulliard who coined the earlier included synonym, A. pseudo-androsaceus. Furthermore, unless Agaricus ericetorum Pers. is considered sanctioned, Art. 7.5 dictates that such illegitimate names are to be "...automatically typified by the type ofthe name which ought to have been adopted under the rules.... " if the author did not designate a type (which Persoon did not do). Therefore, the type of Agaricus ericetorum Pers. (1796) is automatically the type ofAgaricus pseudo­ androsaceus Hull., specifically the central figure on Bulliard's (1786) plate 276. Lange (1981: 693) also dismissed Singer's (1961) typification because, "Singer's notes on the specimen do, however, not indicate that it has the qual ities ofa lectotype." We suppose this refers to the lack of a date. Lange (1981), however, did not offer an alternative type for the name Agaricus ericetorum Pers. Instead he sought to typify Agaricus ericetorum "Fries" as distinct from Agaricus ericetorum Pers. This is illogical if the type method is followed, but understandable given the confusion and broad application ofthe name to several species. Singer (1962) identified Persoon's specimen (L91 0.256-1888, proposed as type ofA. ericetorum Pers.) with the well known, common, palely pigmented species he called Omphalina ericetorum, even while placing the name 0. umbellifera in synonymy. Jorgensen & Ryman (l989a: 140) concurred with the taxonomic identification. The Persoon specimen and the lectotype for both A. pseudo-androsaceus Bull. and A. ericetorum Pers. are considered to be conspecific. 41 However, drawing upon work later reported by Singer & Clemen<;:on (1973), Singer (1975) re-interpreted the type for Omphalina Quelet as being Omphalina umbellifera sensu Quelet, and therefore initiated a schism ofthe treatment ofOmphalina in the literature wherein Agariczls ericetorum "Fr. ex Fr." ended up in his genus Gerronema, but Agarius ericetorum "Pers."was treated as synonymous with both "0. rusticu" and "0. umbellifera (L. ex Fr.) Quel." as well as with "0. pseudoundrosaceous". This contrasted with his earlier assertion the Persoon specimen was typical 0. umbellifera, which all other authors considered to be synonymous with the species he was then calling Gerronema ericetorum. In his 3rd edition ofThe Agaricales in Modern Taxonomy, Singer (1975) proposed retention of the generic name Omphalina Que!. after having moved its type (as he accepted it in the 2nd edition, Singer 1962)to Gerronema(see also Singer 1970). This massive confusion caused by Singer's flip-flop application of the epithet "umbelliferus" and the lack of any single overlapping species between Omphalina sensu Singer (1970, 1975) and Omphalina sensu Bigelow (1970) led Redhead & Weresub (1978) to try to disentangle the synonymy and typifications ofOmphalina. Together these publications precipitated the discussion by Lange (1981).

In the absence ofconserved types for any ofthe species names, despite long debate, the typifications must again be reconsidered in light of revised wording of the Code (Greuter et aI., 2000). Art. 7.8 currently states, "The typification ofnames adopted in one ofthe works specified in Art. 13.1 (d), and thereby sanctioned (Art. 15), may be effected in the light ofanything associated with the name in that work." Lange (1981 : 695) sought to neotypify Agaricus ericetorum "Fr. ex Fr." with Lundell & Nannfeldt, Fungi exsicc. Suec. No. 1753 (C). J0rgensen & Ryman (1989a) noted that the specimen could not be a neotype because original material existed, namely illustrations cited by Fries (1821). They later (J0rgensen & Ryman 1994: 254) selected Tab. 1015, ut "Agaricus niveus" (Vahl 1790), as lectotype, formally epitypifying it by a specimen Lundell & Nannfeldt, Fungi exsicc. Suec. No. 1753 in herbarium UPS (rather than in C as was suggested by Lange 1981). The species represented by this specimen (to judge by all accounts on portions in other herbaria, e.g., C & DAOM), is conspecific with the species represented by the lectotype ofA. ericetorum Pers. They are the same if 42 the type method is followed. If they are the same, it cannot be argued that there is both an Agaricus ericetorum Pers. and an Agaricus ericetorum "Fr." Ifthey are conspecific, and the one name is based on the other name, logically and nomenclaturally they must have the same lectotype. Fries (1821: 509) himself attributed the name to Persoon. Redhead & Kuyper (1987: 329) had selected the earliest lectorype for Agaricus ericetorum of Fries (1821) as well as ofAgaricus ericetorum Pers. and A. pseudo-androsaceus Bull. This predates the selection of lectotype by J0rgensen & Ryman (1989a). The selection of an epitype by J0rgensen & Ryman (1994), however, is uncontested. It clearly is applicable to the name Agaricus ericetorum Pers.: Fr.

Controversy continued to dog the typification ofAgaricus umbelliferus L. and Omphalina Quelet (J0rgensen & Ryman 1989a, b, Lange 1981, Redhead 1993, Redhead & Kuyper 1987, 1988, 1993, Singer 1986). We note that subsequent changes in the Code now allow different interpretations from those allowed in the 1980's. In particular, Art. 13.(d) does not disallow lichenized fungi from having sanctioned names, and as noted above Art. 7.8 state, "The typification ofnames adopted in one ofthe works specified in Art. 13.1 (d), and thereby sanctioned (Art. IS), may be effected in the light ofanything associated with the name in that work." This would allow for the rejection ofthe proposed lectotype ofAgaricus umbelliferus L. by Redhead & Kuyper (1987, 1993), by the fact that Fries (1821) placed that proposed "lectotype" in another species. This in turn allows us to accept the lectotypification proposed by J0rgensen & Ryman (1989a), namely an unpublished original illustration by Rudbeck in the year 1695, apparently available to Linnaeus, which has been posthumously reproduced and published (Rudbeck 1987). A change in the wording ofArt. 9.2 (Note 2) regarding "original material" nullifies the nomenclatural argument put forward by Redhead & Kuyper (1993) against this typification. We also accept the same epitype as that for the name Agaricus ericetorum Pers.: Fr., proposed by J0rgensen & Ryman (1994), i.e., Lundell & Nannfeldt, Fungi exsicc. Suec. No. 1753, herb. UPS. These lecto- and epi­ typifications effectively makeAgaricus umbelliferus L.: Fr. synonymous with Agaricus ericetorum Pers.: Fr. It is hoped that this compromise resolves and ends the 250-year old controversy over these names. Among sanctioned names, Agaricus umbelliferus has priority (Gams & 43 Kuyper 1984) and its lichenized status no longer nomenclaturally affects its availability.

L. velutina (Quel.) comb. nov. Basionym: Omphalia velutina Quet., C.R. Ass. frans:. Av. Sci. (Grenoble, 1885) 14: 445. 1886.

Comments: The following names represent taxa in a species complex, characterized by greyish basidiomata with incrusting pigments, clampless hyphae, velutinous stipes, elongated basidiospores, and a thallus consisting of numerous loose globose Phytoconis-like "goniocysts (cf. Serusiaux 1985)" linked by characteristically very narrow hyphae (1-)2(-3),um diam (see Redhead & Kuyper 1987). They are considered to be synonymous at a species level, although a phylogenetic study by Lutzoni (1997) suggests that more than one species might be recognized molecularly in this complex, hence our recognition ofL. grisella:

?= Omphalina oreades Singer and its nom. nov. Clitocybe albimontana Bigelow (see discussion in Redhead & Kuyper 1987).

?= Omphalina pararustica Clemens:on (fide Kuyper 1995). Note: said to differ from L. velutina by presence of4-spored basidia.

?= Clitocybe borealis Bigelow and its nom. nov. Clitocybe kemptonii Bigelow [Holotype at MICH examined by SAR and thalli confirmed; V. Wells & P. Kempton #2298, 13 Jul 1966, Eklutina L., Alaska, USA; also examined were specimens cited by Bigelow (1985), i.e. Wells & Kempton #2410, Alaska, conspecific; A.H. Smith # 55960, California, conspecific; and a specimen identified as C. kemptonii by H.E. Bigelow at MICH but not cited by him, i.e. A.H. Smith #80399, 31 Aug 1971, alpine meadow, Davos-Platz, Switzerland]. Excluded from "e. kemptonii": Wells & Kempton #2409, Alaska - a clamped, nonlichenized omphalinoid Arrhenia cited by Bigelow (1985) as C. kemptonii.

?= Omphalina defibulata Singer 1952 [Isotype at MICH examined by SAR and thalli confirmed; R. Singer M355, 10 Feb 1950, Tierra 44 del Fuego, Rio Grande, Estancia, Nueva, Argentina]

?= Clitocybe solumophila Bigelow 1983 [Holotype at MICH examined by SAR and thalli confirmed; A.H. Smith #850 1,6 Nov 1937, Siskiyou Natl. Forest, Siskiyou, California, USA]

?= Clitocybepayettensis Bigelow 1983 [Holotype at MICH examined by SAR and thall i confirmed; A.H. Sm ith #44296, 28 June 1954, Lake Fork Creek, Valley Co., Idaho, USA]

'1= Omphalina meridionalis Contu & La Rocca, Fungi non delineati 9: 32-33. 1999. [Holotype re-examined and reinterpreted as a clampless species by Barrasa & Esteve-Raventos 2000. Confirmation of the clampless state was made on a later collection kindly collected, identified and supplied by Marco Contu for deposit in DAOM (#229481), Italy, Sardinia, Trinita, d'Agultu, loc. Vignola, 18 Feb. 2001. However, this sample has zebra-striped, finely incrusted pigments contrary to the smoothly pigmented hyphae reported by Barrasa & Esteve­ Raventos (2000). It also has the narrower thallic hyphae typical for the L. velutina complex. It remains debatable as whether or not and how to distinguish this taxon from L. velutina or other named 'species' in the complex. The other Lichenomphalia species (L. umbellifera, L. alpina, L. hudwniana) all exhibit variations in pigmentation throughout their geographic ranges (Redhead & Kuyper 1987).]

Note: The selection ofnames and typification ofLichenomphalia listed above bring into conformity the species epithets adopted for Omphalina by M. Lange in Nordic Macromycetes Vol. 2 (Hansen & Knudsen 1992) and by Senn-Irlet et al. (1990). We hesitate to transfer to Lichenomphalia the steriIe lichen, Omphalinafoliacea Jorgensen (1989), which remarkably was described as an Omphalina despite the lack of known basidiomata (or ascomata), or any proof, other than suspicion, that it represents a Basidiomycete.

Arrhenia Fries, Summa Veget. p. 312. 1849. = Boehmia Raddi 1807 nom. rej. (Greuter et al. 2000) 45

= Corniola Gray 1821 non Corniola Adans. 1763 = Leptotus P. Karst. 1879 = Leptoglossum P. Karst. 1879 = DietyoIus Quel. 1886 = Geotus Pilat & Svrcek 1953 = Phaeotellus Kuhner & Lamoure 1972 = Arrhenia subg. Phaeotellus (Kuhner & Lamoure) Kuhner 1980 = Clitoeybe subg. Atroviridae Bigelow 1982 = Omphalina sect. Atroviridae (Bigelow) Redhead 1986 = Leptoglossum subg. Phaeotellus (Kuhner & Lamoure) Singer 1986

Previously transferred and accepted species ofcyphelloid, nutant,or cantharelloid taxa:

A. aeerosa (Fr.) Kuhner (1980: 893 & 992) (type for Phaeotellus Kuhner & Lamoure 1972) ?= A. latispora (Favre) Bon & Courtecuisse (1987: 37) A. alnetorum (Singer) Redhead (1984: 875) Note: correction ofepithet, genitive plural of alnetum, i.e. alder wood), supplied by I.A. Nannfeldt (pers. comm. 1984). A. auriscalpium (Fr.) Fr. (1849: ) (lectotype for Arrhenia Fr. 1849) A. australis (Clel.) Grgurinovic (1997: 318) A.jissa (Leys.) Redhead (1984: 875) A. griseopallida (Desm.) Watling (1988: 553) A. lobata (Pers.: Fr.) Kuhner & Lamoure ex Redhead (1984: 871) (lectotype for Dictyolus Quel. 1886) A. retiruga (Bull.: Fr.) Redhead (1984: 873) (lectotype for Leptotus P. Karst. 1879) A. riekenii (Hora) Watling (1988: 553) A. salina (Heiland) Bon & Courtecuisse (\ 987: 37) [superfl. comb. by Gulden 1988] A. spathulata (Fr.) Redhead (1984: 876) (= Cantharellus muscigenus (Bull.: Fr.) Fr. lectotype for Leptoglossum P. Karst. 1879; non Arrhenia museigena (Pers.: Fr.) Que!. 1888 = Cyphellostereum laeve (Fr.) Reid, see Redhead 1984: 876) 46 Required new combinations:

Arrhenia andina (Corner) comb. nov., Basionym: Leptoglossum andinum Corner, Monogr. Cantharelloid Fungi, p. 140. 1966. Arrhenia antarctica (Singer) comb. nov. Basionym: Omphalina antarctica Singer, Beih. Sydowia 1: 16. 1956. Arrhenia baeospora (Singer) comb. nov., Basionym: Omphalina baeo/!'pora Singer, Z. Pilzk. 43: 120. 1977. Arrhenia chilensis (Mont.) comb. nov. Basionym: Agaricus chi/ensis Mont., Ann. Sc. Nat. II, 8: 368. 1837. Arrhenia c111orocyanea (Pat.) comb. nov. Basionym: Agaricus chlorocyaneus Pat., Tab. anal. Fung. 4: 145. 1885. Comments: The taxonomic concept adopted here is that of Redhead (1986) under the name Omphalina viridis (Hornem.) Kuyper (= Agaricus ericetorum var. viridis Homem., incorrectly listed as "Agaricus umbelliferus var. viridis Homem." by Kuyper 1995 as a synonym of Omphalina chlorocyanea (Pat.) Singer). Clitocybe atroviridis Bigelow (1982) was treated as a synonym by both Redhead (1986) and Kuyper (1995). Bigelow (1985: 413-414), defended publication of his new species, but agreed that European collections that he had earlier reported to lack clamp connections (Bigelow 1982), in fact had clamp connections. He then noted that a North American collection (A.H. Smith 70705, MICH) representing "Omphalina umbellifera var. viridis" lacked clamp connections. However, that collection was later examined by SAR and redetermined as Chrysomphalina grossula (Pers.) Norvell et aI., a clampless, much paler species lacking bluish green coloration (Norvell et al. 1994). Nonetheless, Bigelow'S (1982) name, Clitocybe atroviridis, is the only uncontroversial name linked to the vernal species represented by extant specimens because ofthe availability ofBigelow's type specimen. Having established that reports of a c1ampless species similar to the currently recognized one were either false or based upon a wholly different species and genus, there remains only the presence of hairs on the stipes, as depicted by Patouillard (1885) as a barrier to acceptance of Patouillard's earlier name. Specimens identified as A. chlorocyaneus that might have served as type, are absent from Patouillard's herbarium in the Farlow herbarium, as confirmed by the latc H.E. Bigelow (pers. comm. to SAR), and by L.L. Norvell & D. Pfister more recently (pers. com. to SAR). Almost all collections, ofthe 47 bluish green omphalinoid taxon cited by Redhead (1986) and Bigelow (1982) have glabrous stipes, but on some there are projecting hyphaI ends, sometimes drawn into wisps (e.g., DAOM 188208). Environmental conditions, if not genetic variability, may well explain the occasional occurrence ofminutely pubescent stipes. In the absence ofthe discovery ofanother vcrnal bluish green species in France, and the predominantly spring fruitings of the species as we know it in both Europe and North America, we adopt here the epithet published by Patouillard.

Arr/lenia epichysium (Pers.: Fr.) comb. nov. Basionym: Agaricus epiehysium Pers. Syn. Meth. Fung. p. 462. 1801. [Agaricus epiehysium Pers.: Fr., Syst. Mycol. I: 169. 1821.] Arr/lenia etegans (Pers.) comb. nov. Basionym: Merulius elegans Pers., Syn. Meth. Fung. p. 481. 1801. Arrhenia hohensis (A.H. Smith) comb. nov. Basionym: Omphalia hohensis A.H. Smith, Contr. Univ. Mich. Herb. 5: 27. 1941. Arrhenia tundellii (Pilat) comb. nov., Basionym: Omphalia lundellii Pilat, Friesii 5: 26-28. 1954. Arrhenia oblltra (Favre) comb. nov. Basionym: Omphalia ohatra Favre, Les Champignons superieurs de la zone alpine du Parc National Suisse. Resultats des recherches scientifiques entreprisesau ParcNational suisse. 5 (N.S.): 46,199.1955 [see Lamoure 1975]. Arrhenill obscumta (D. Reid) comb. nov., Basionym: Omphalina obseurata D. Reid, Trans. Brit. Mycol. Soc. 41: 419. 1958. [Note: = Clitoeybe atrobrunnea Bigelow 1985, nom. nov.]. Arrhenia omnivora (Agerer) comb. nov. Basionym: Leptoglossum omnivorum Agerer, Trans. Brit. Mycol. Soc. 82: 184. 1984. [Holotype at K examined by SAR; the assumption is made here that the subtle membranal pigments are indications ofaffinities with other Arrhenia species, rather than with Rimbaehia]. Arrheniaonisca (Fr.: Fr.) comb. nov. Basionym: Agaricus aniseus Fr., Obs. Mycol. 2: 209. 1818. [Agaricus oniseus Fr.: Fr., Syst. mycol. 1: 172. 1821]. Arrhenia parvivelutina (Ch~ment;on & Irlet) comb. nov., Basionym: Omphalina parvivelutina Clemen90n & Irlet, Schw. Z. Pilzk. 1982A Sondemummer 123 (Mycologia Helvetica): 15. 1982. Arr/lenia pauxillll (Clemen~on) comb. nov., Basionym: Omphalina 48 pauxilla Clemenlj:on, Z. Mykol. 48: 213. 1982. Arrhenia peltigerina (Peck) comb. nov. Basionym: Agaricus peltigerinus Peck, Rep. New York State Mus. (for 1876) 30: 38. 1878. Arrhenia phi/onotis (Lasch) comb. nov. Basionym: Agaricus philonotis Lasch, Linnaea 3: 394.1828. Arrhenia pubescentipes (Bigelow) comb. nov. Basionym: ClitoGybe pubescentipes Bigelow, Beih. Nova Hedw. 81: 389. 1985. Arrhenia rainierensis (Bigelow) comb. nov. Basionym: ClitoGybe rainierensis Bigelow, Beih. Nova Hedw. 81: 390. 1985. Arrhenia rigidipes (Lamoure) comb. nov. Basionym: Omphalina rigidipes Lamoure, Arctic & Alpine Mycol. (First IntI. Symp. Arcto-Alpine Mycology) p. 207. 1982. Arrhenia rustica (Fr.) comb. nov. Basionym: Agaricus rusticus Fr., Epicr. Syst. Mycol. p. 126, 1838. Note: see J0rgensen & Ryman 1989b: 141 for typification. Arrhenia sphaerospora (Lamoure) comb. nov. Basionym: Omphalina sphaerospora Lamoure, Trav. Sci. Parc Natl. Vanoise 6: 164. 1975. Arrhenia sphagnicola (Berk.) comb. nov. Basionym: Agaricus sphagnicola Berk., British Fungi, English Flora 5(2): 67. 1836. (Note: see Redhead & Kuyper 1987 for comments on types). Arrhenia stercoraria (Barrasa, Esteve-Rav. & Sanchez) comb. nov. Basionym: Glabrocyphella stercoraria Barrasa, Esteve-Rav. & Sanchez MycoI.Res.102: 1265.1998. Arrhenia subglobispora (Moreno, Heykoop & Horak) comb. nov. Basionym: Omphalina subglobispora Moreno, Heykoop & Horak, Mycotaxon 77: 366.2001. Arrhenia suhobscura (Singer) comb. nov. Basionym: Omphalina subobscura Singer, Flora Neotropica 3: 21. 1970. Arrhenia !mbumbratilis (Singer) comb. nov. Basionym: Omphalina subumbratilis Singer, Flora Neotropica 3: 23. 1970. Arrhenia trigono!ipora (Lamoure) comb. nov. Basionym: Omphalina trigonospora Lamoure, Trav. Sci. Parc Natl. Vanoise 6: 157. 1975. Arrhenia umhratilis (Fr.: Fr.) comb. nov. Basionym: Agaricus umbratilis Fr.:Fr. Syst. Mycol. I: 157. 1821. Arrhenia velutipes (P.O. Orton) comb. nov. Basionym: Omphalina 49 velutipes P.O. Orton, Trans. Brit. Mycol. Soc. 43: 337. 1960. Arrhenia vi;'idimammata (Pilat) comb. nov. Basionym: Omphalia viridimammata Pilat, Friesii 5: 25-26. 1954. Arrhenia volkertii (Murr.) comb. nov. Basionym: Omphalia volkertii Murrill, Mycologia 3: 98. 1911. [see Bigelow 1985 for type study].

Acknowledgements This work was supported in part by NSF grants DEB-0076023 and DEB­ 9708035. SAR thanks the past and present curators ofthe Michigan herbarium (MICH) and Kew Gardens herbarium (K) for the loan of type materials, and Marco Contu (Italy) for supplying a specimen of Omeridionalis. We thank Keith Seifert, Robert Shoemaker and Lorelei L. Norvell for presubmission reviews. SAR also acknowledges the assistance of the late H.E. Bigelow and J.A. Nannfeldt on nomenclature and/or taxonomy.

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